Rondeletieae (Rubiaceae): Part I (Rustia, Tresanthera ... - CNCFlora

10 Flora Neotropica
mineeae, but had reservations about including the PHYLOGENETIC RELATIONSHIPS
group in the Catesbaeeae (sensu Hooker). The genera
listed under the Condamineeae by Robbrecht
(1993) were Chimarrhis, Condaminea, Dioicodendron,
Flexanthera [= Simira], Kerianthera [Isertieae],
Parachimarrhis, Picardaea, Pinckneya, Pogonopus,
Rustia (incl. Stomandra), Tresanthera, and tentatively
Kajewskiella Merr. & Perry, with Pseudomussaenda
Wernham transferred to the Isertieae. Robbrecht
(1993) made several changes in the genera that he
previously included in the Rondeletieae (Robbrecht,
1988). Following Lorence (1991), Robbrecht (1993)
treated Arachnothryx, Rogiera, and Javorkaea as synonyms
of Rondeletia, and also added Mazaea Krug
& Urb. He maintained the Sipaneeae (to which he
added Neobertiera Wernham) and separated the
Simireae (here included, and proposed to be closely
related to Parachimarrhis) from the Rondeletieae.
At the Second International Rubiaceae Conference
(13-15 September 1995, Meise, Belgium), Bremer
(1996) presented a second phylogenetic analysis using
rbcL molecular sequences that supported her previous
results (Bremer, 1995) concerning the monophyletic
origin of the Rubiaceae and their subdivision
into three main lineages. Delprete (1995a, 1996d)
presented a second set of tribal-level phylogenetic
analyses using morphological data, from 170 species
of 44 genera, including all Condamineeae and Catesbaeeae
sensu Hooker and selected Chiococceae, Rondeletieae,
Cinchoneae, and Coffeae. According to the
results of these analyses (Delprete, 1995a, 1996d), the
two subtribes Condamineinae and Pinckneyinae were
transferred into a broader, tentative delimitation of the
Rondeletieae, which is adopted, with few modifications,
in the present treatment. He also included the
subtribe Portlandiinae (Condamineeae sensu Hooker,
1873) in the Catesbaeeae. The tribe Catesbaeeae
(sensu Hooker, 1873) has nomenclatural priority over
the subtribe Portlandiinae (also, Portlandieae Baillon
is a nomen illegitimum because it includes Rondeletia);
therefore, the newly emended tribe should be called
"Catesbaeeae J. D. Hooker emend. Delprete" (Delprete,
1996d). An abbreviated version of these analyses is
presented in Phylogenetic Relationships, below.
Classification at subfamilial and tribal levels in the
Rubiaceae has been debated since its establishment,
and up to the present day no overall consensus among
workers has been reached. As certain taxonomic characters
are carefully studied, or new ones are added to
phylogenetic analyses, the macrosystematics of the
Rubiaceae have been reexamined and modified accordingly.
The number of recognized subfamilies within the
Rubiaceae varied according to systematists' interpretation
of the usefulness and taxonomic importance of
certain characters. Using similar data sets, Verdcourt
(1958) proposed three subfamilies (Rubioideae,
Cinchonoideae, and Guettardoideae) whereas
Bremekamp (1966) divided the Rubiaceae into the
largest number of subfamilies ever proposed, eight:
Cinchonoideae, Urophylloideae, Pomazotoideae,
Gleasonioideae, Guettardoideae, Ixoroideae, Rubioideae,
and Hillioideae. The most recent comprehensive
classification using morphological, anatomical,
and cytological data was proposed by Robbrecht
(1988, 1993), who recognized four subfamilies
(Cinchonoideae, Rubioideae, Antirheoideae, and
Ixoroideae) and 44 tribes (a compromise between
Verdcourt's and Bremekamp's classifications).
Bremer (Bremer et al., 1995; Bremer, 1996) presented
phylogenetic analyses using rbcL molecular sequences
showing evidence for recognition of only
three subfamilies (merging the Antirheoideae within
the Ixoroideae s.l.), which was later supported by
trnL-F (Rova et al., 1997) and rps 16 (Andersson,
unpubl. data) molecular sequences.
The phylogenetic analyses using molecular and
morphological data presented by Bremer (1992),
Bremer and Eriksson (1992), and Bremer and Struwe
(1992) supported Bremekamp's and Verdcourt's opinions
about the ambiguous usefulness of certain flower
and fruit characters. In my opinion, fruit and flower
characters have often been superficially or erroneously
interpreted in trying to detect phylogenetic relationships.
In order to avoid the repetition of erroneous
observations, morphological characters and characters
states should be coded after direct observation of fresh
Rova et al. (1997) presented a phylogenetic analysis
using trnL molecular sequences of more than 100
species representative of the most tribes in
Robbrecht's (1988, 1993) classification. These analyses
supported the rbcL phylogenies of Bremer et al.
(1995) and Bremer (1996), and rpsl6 phylogenies of
Andersson (in progress, unpubl. data) in dividing the
Rubiaceae in three main groups: Cinchonoideae s.str.,
Ixoroideae s.l., and Rubioideae.
or preserved material, instead of being reported from
previous literature. The taxonomic significance of
characters such as number of ovules per locule, placentation,
mesocarp fleshiness, seed insertion and
position within the fruit, dioecy, heterostyly, and corolla
aestivation should be evaluated with accurate
anatomical and morphological studies, and compared
with phylogenies obtained with alternative sets of data
(i.e., molecular, biochemical, cytological data).